Embryogenesis of the dipluran Lepidocampa weberi Oudemans (hexapoda: diplura, campodeidae): formation of dorsal organ and related phenomena

The developmental changes of embryonic membranes of a dipluran Lepidocampa weberi, with special reference to dorsal organ formation, are described in detail by light, scanning, and transmission electron microscopies. Newly differentiated germ band and serosa secrete the blastodermic cuticle at the entire egg surface beneath the chorion. Soon after, the serosal cells start to move dorsad. All the serosal cells finally concentrate at the dorsal side of the egg and form the dorsal organ. During their concentration, the serosal cells attenuate their cytoplasm to form filaments. The extensive area from which the serosa has receded is occupied by a second embryonic membrane, the amnion, which originates from the embryonic margin. The embryo and newly emerged amnion then secrete three fine cuticular layers, "cuticular lamellae I, II, and III," above which the filaments of the (developing) dorsal organ are situated. With the progression of definitive dorsal closure, the amnion reduces its extension, the dorsal organ is incorporated into the body cavity of the embryo, and the amnion and dorsal organ finally degenerate.The dorsal organ of diplurans is formed by the concentration of whole serosal cells, while that of collembolans is formed by the direct differentiation of a part of serosal cells. However, the dorsal organs of diplurans and collembolans closely resemble each other in major aspects, including that of ultrastructural features, and there is no doubt regarding their homology. The amnion, which has been regarded as being a characteristic of Ectognatha, also develops in the Diplura. This might suggest a closer affinity between the Diplura and Ectognatha than previously believed.     

Blastodermic cuticles of a springtail,Tomocerus ishibashii Yosii (Collembola : Tomoceridae)

The formation and structure of the blastodermic cuticles of a springtail, Tomocerus ishibashii Yosii (Collembola : Tomoceridae) are described together with the change of egg membrane. The blastodermic cuticles of the Collembola are 2-layered, and formed in the early stages of the embryonic development, preceding the differentiation of germ band. The first blastodermic cuticle is thicker (about 0.8-1.5 μm in thickness) and its surface is provided with complex structures, whereas the second one is thinner (about 0.2-0.4 μm in thickness) and smooth. About 3 days after oviposition, the chorion (about 2 μm in thickness) splits into 2 and the first blastodermic cuticle, provided with many projections and 4 large spines appear on the surface of the egg. Three types of projections are distinguished: button-, cone- and seta-like structures. The halves of the ruptured chorion are attached to the first blastodermic cuticle on both sides below the spines, and no projections are found in the regions concealed by the ruptured chorion. The projections of the first blastodermic cuticle are formed by cellular protrusions of the blastoderm. The conspicuous large spines on the first blastodermic cuticle are formed by the evaginations of the blastoderm. Tendrils of the primary dorsal organ run between the first and second blastodermic cuticles.     

Ultrastructure of the embryonic dorsal organ of Orchestia cavimana (crustacea, amphipoda); with a note on localization of chloride and on the change in calcium-deposition before the embryonic moult

The transitory dorsal organ of Orchestia cavimana appears simultaneously with the development of the germ layer and is gradually reduced during the last 2-3 days of embryonic development. It represents the only direct connection of the embryo with the chorion or-after the embryonic moult-with the embryonic envelope. The shape is hemispherical and it consists of about 50 bottle-shaped cells, arranged radially around a centre. This centre is filled with different kinds of extracellular material which forms a central plug apically and a central cone below it. The bottle-shaped cells taper apically. The neck region of these cells is characterized by numerous microvilli which project into the intercellular space. This space is filled with an electron dense substance and is in contact with the central cone. In the basal neck region numerous mitochondria are associated with the microvilli. The high density of mitochondria is characteristic for the nuclear region. The cytoplasm of the basal region below the nucleus contains numerous calcium granules. Evidence for the concentration of chloride in the apical dorsal organ is shown. Before the embryonic moult and during the duplication of the egg-volume the number of calcium granules in the dorsal organ and the integument is reduced. Simultaneously calcium granules appear in the now visible periembryonic space. This suggests that part of the calcium is shifted into this space. The function of the dorsal organ is discussed. Besides the probable main function-transport activity of ions-its role before and during embryonic moult and its part in the utilisation of yolk are discussed.     

Embryogenesis of the honeybee apis mellifera l. (hymenoptera : apidae): An sem study

Our SEM study of honeybee, Apis mellifera (Hymenoptera : Apidae), embryogenesis is based on embryos fixed at 1 hr intervals from oviposition to hatching. Embryos of equal age showed little variation, so that staging could be based on developmental age. Our data confirm many earlier light microscopical observations, but are at variance with some others. The cytoplasmic connections between the future blastoderm cells and the central yolk system are severed only at the onset of gastrulation. The serosa derives from cells which immigrate into the dorsal strip and then join up to form a pre-serosa bordering the germ band rims. When the serosa has detached, the amnion grows out from the germ band margins and serves as a provisional dorsal epithelium right from the beginning. Germ band segmentation is followed by the transient regression of every second transverse groove (double segment pattern). The germ band flanks grow dorsally and replace the amnion a few hr before hatching (dorsal closure). The tracheal openings which form half-way between segment borders are closed temporarily by the embryonic cuticle; similar openings above the labial buds contribute to the tentorium rather than the tracheal system. Most head appendages retain bud character until long after hatching. The events observed in the SEM are linked in a diagram to the stage series based on living embryos.     

Embryonic integument and "molts" in Manduca sexta (Insecta,Lepidoptera)

In Manduca sexta the germ band is formed 12 h post-oviposition (p.o.) (=10% development completed) and is located above the yolk at the egg surface. The cells show a polar organization. They are engaged in the uptake and degradation of yolk globules, pinched off from the yolk cells. This process can be observed in the integumental cells during the first growth phase of the embryo that lasts until "katatrepsis," an embryonic movement that takes place at 40% development completed. At 37% development completed, the ectoderm deposits a thin membrane at its apical surface, the first embryonic membrane, which detaches immediately before katatrepsis. The second period of embryonic growth--from katatrepsis to 84 h p.o. (70% development completed)--starts with the deposition of a second embryonic membrane that is somewhat thicker than the first one and shows a trilaminar, cuticulin-like structure. Whereas the apical cell surface is largely smooth during the deposition of the first embryonic membrane, it forms microvilli during deposition of the second one. At the same time, uptake of formed yolk material ceases and the epidermal cells now contain clusters of mitochondria below the apical surface. Rough endoplasmic reticulum (RER) increases in the perinuclear region. The second embryonic membrane detaches about 63 h p.o. At 69 h p.o., a new generation of microvilli forms and islands of a typical cuticulin layer indicate the onset of the deposition of the larval cuticle. The third growth phase is characterized by a steady increase in the embryo length, the deposition of the larval procuticle, and by cuticular tanning at about 100 h p.o. Beginning at that stage, electron-lucent vesicles aggregate below the epidermal surface and are apparently released below the larval cuticle. Manduca sexta is the first holometabolous insect in which the deposition of embryonic membranes and cuticles has been examined by electron microscopy. In correspondence with hemimetabolous insects, the embryo of M. sexta secretes three covers at approximately the same developmental stage. A marked difference: the second embryonic cover, which in Hemimetabola clearly exhibits a cuticular organization, has instead a membranous, cuticulin-like structure. We see the difference as the result of an evolutionary reductional process promoted by the redundancy of embryonic covers in the egg shell. Embryonic "molts" also occur in noninsect arthropods; their phylogenetical aspects are discussed.     

The serosa of Manduca sexta (Insecta, Lepidoptera): ontogeny, secretory activity, structural changes, and functional considerations

In Manduca sexta, the blastoderm forms successively and becomes immediately cellularized as the cleavage energids reach the surface of the oocyte. Presumptive serosal cells are large and contain 2 or 4 large polyploid nuclei; presumptive embryonic cells are small and mononuclear. All parts of the blastoderm participate in the uptake and digestion of yolk material. About 10 h post-oviposition, the blastoderm breaks at the amnioserosal fold and the extraembryonic part closes above the germ band and constitutes the serosa (12 h post-oviposition, i.e. 10% development completed). At once, the serosa starts to secrete a cuticle consisting of an epi- and a lamellated endocuticle. Detachment of the serosal cuticle, 22h post-oviposition, is reminiscent of apolysis of larval cuticle. Thereafter, the serosa deposits a membranous structure, the serosal membrane. The sercretory process lasts from 23h to 44h post-oviposition. At first a fine granular layer, then an amorphous, spongy-like, fibrillar layer is secreted via microvilli. This persisting membrane is tough, rubbery and very elastic. It may serve to bolster the serosa during katatrepsis (48h post-oviposition) and later embryonic movements. After detachment of the serosal membrane, 44h post-oviposition, a distinct subcellular reorganization of the serosa takes place. The nuclei become still larger and more irregular. Uptake of yolk granules, but not of lipid droplets, ceases, although interaction of serosa and yolk cells are intense. Serosal cells include many mitochondria, large areas of rER, besides some sER, increasing amounts of lysosomal bodies and prominent Golgi complexes. Most conspicuous is the assembly of spindle-shaped, electron-lucent vesicles below the apical surface. These vesicles may contain metabolic products which are released into the peripheral space. The studies show that the serosa assumes changing functions during embryogenesis: digestion of yolk substances, synthesis of a serosal cuticle and a serosal membrane, which may have a protective function, and excretion.     

Ecdysteroid Titres and Cuticle Depositions in Embryos of the Dipteran Calliphora erythrocephala

Summary

Before the shortening of the germ band, embryos of Calliphora erythrocephala secrete an electron dense layer which resembles a thin atypical cuticle. After completion of the dorsal closure, a second, typical, cuticle is deposited. It is characterized by a thick procuticle and by the presence of hooked setae. This cuticle develops into the larval cuticle of the first instar.

We did not find in newly-laid eggs of Calliphora detectable amounts of either free or hydrolysable conjugated ecdysteroids. A marked rise in ecdysteroid concentrations, essentially attributable to free ecdysone and 20- hydroxyecdysone, occurs shortly before the deposition of the typical cuticle. After a transient decrease, the ecdysteroid titre rises again before hatching.

When eggs are mid-ligatured at blastula or early gastrula stages, the posterior embryonic half is able to build up a typical cuticle with hooked setae. This cuticulogenesis is therefore not dependent on the presence of the embryonic ring glands.     

Stage-dependent strategies of host invasion in the egg-larval parasitoid Chelonus inanitus

Chelonus inanitus (Braconidae) is a solitary egg-larval parasitoid which lays its eggs into eggs of Spodoptera littoralis (Noctuidae); the parasitoid larva then develops in the haemocoel of the host larva. Host embryonic development lasts approx. 3.5 days while parasitoid embryonic development lasts approx. 16 h. All stages of host eggs can be successfully parasitized, and we show here that either the parasitoid larva or the wasp assures that the larva eventually is located in the host's haemocoel. (1) When freshly laid eggs, up to almost 1-day-old, are parasitized, the parasitoid hatches while still in the yolk and enters the host either after waiting or immediately through the dorsal opening. (2) When 1-2-day-old eggs are parasitized, the host embryo has accomplished final dorsal closure and is covered by an embryonic cuticle when the parasitoid hatches; in this case the parasitoid larva bores with its moving abdominal tip into the host. (3) When 2.5-3.5-day-old eggs are parasitized, the wasp oviposits directly into the haemocoel of the host embryo; from day 2 to 2.5 the embryo is still very small and the wasps, after probing, often restrain from oviposition for a few hours.     

Embryonic development of the jumping bristletail Pedetonutus unimaculatus Machida,with special reference to embryonic membranes (Hexapoda: Microcoryphia,Machilidae)

In the machilid Pedetonutus unimaculatus, a germ disc is formed by the aggregation and proliferation of cells within a broadly defined embryonic area. Cells adjacent to the embryonic area form the serosal fold that grows beneath the embryo. Then the embryonic margin is extended to form a cell layer or amnion that lies between the embryo and serosal fold. Thus, an amnioserosal fold is formed by the addition of the amnion to the serosal fold. Serosal cells cover the entire surface of the egg and begin to secrete a serosal cuticle. Soon the amnioserosal fold is withdrawn, and the embryo is exposed to the egg surface. The spreading amnion replaces the serosal cells that finally degenerate through the formation of a secondary dorsal organ. In the areas of amnion anterior and lateral to the embryo, yolk folds form and encompass the embryo. The amnion is a provisional dorsal closure and never participates in the formation of the definitive one. The amnioserosal fold of the Microcoryphia appears to have the functional role of secreting a serosal cuticle beneath the embryo. This fold of the Microcoryphia may be regarded as an ancestral form of the amnioserosal folds of the Thysanura-Pterygota. the yolk folds may appear to be passive transformation of the yolk mass linked to positioning of the growing embryo within the egg. There is no evidence that the yolk folds and the cavity appearing between them in the Microcoryphia are homologous to the amnioserosal fold and amniotic cavity in the Thysanura-Pterygota. The yolk folds appear to be one of the embryological autapomorphies in the Microcoryphia. © 1994 Wiley-Liss, Inc.     

Cells released in vitro from the embryonic yolk sac of the stick insect carausius morosus (BR.) (PHASMATODEA : HETERONEMIIDAE) may include embryonic hemocytes

The embryonic yolk sac and the adult dorsal vessel of the stick insect Carausius morosus (Br.) (Phasmatodea : Heteronemiidae) were shown to release a number of cells that appear morphologically similar to circulating adult hemocytes. Like adult hemocytes, these cells reacted positively when tested for both phenoloxidase activity and a monoclonal antibody specifically raised against a vitellin polypeptide. Based on this evidence, it is suggested that yolk sac-released cells behave as potential embryonic hemocytes. A model is thus proposed whereby the yolk sac might host a number of hemopoietic stem cells on their way to the dorsal vessel, and in so doing, it may temporally act as an embryonic hemopoietic organ.     

Ultrastructure of alimentary tract formation in embryos of two insect species: Melasoma saliceti and Chrysolina pardalina (Coleoptera, Chrysomelidae)

Embryogenesis of the alimentary tract in two chrysomelid species (Chrysolina pardalina and Melasoma saliceti) is described. The embryonic development of both species lasts 7days at room temperature. Stomodaeum and proctodaeum invaginate at the anterior and posterior ends of the germ band. Together with the ectodermal tissue the endoderm cells also enter into the embryo. The anterior and posterior parts of the alimentary tract wedge into the yolk in the form of conical structures. The endodermal cells remain at the yolk surface and start migration over the yolk mass as two lateral bands of cells. The endoderm is always accompanied by mesoderm. On the fifth day of development the endodermal cells together with the mesoderm layer spread over the ventral and dorsal sides of the yolk mass and form the single layered primordium of the midgut epithelium. On the sixth day of development a basal lamina appears between the endoderm and the mesoderm cells and differentiation of both tissues starts. The endodermal epithelium cells change shape from flat to cuboidal and eventually into columnar. Mesoderm cells differentiate into muscle and tracheae. On the 7thday of development stomodaeum and proctodaeum become lined with cuticle and the midgut becomes covered with microvilli. The yolk cells populating the yolk mass do not contribute to midgut formation in the species studied.     

Ejaculatory duct of the migratory grasshopper,Melanoplus sanguinipes (fabr.) (Orthoptera : Acrididae): A histological and ultrastructural analysis

The ejaculatory duct of the migratory grasshopper (Melanoplus sanguinipes [Fabr.]) (Orthoptera : Acrididae) is divisible into 3 regions: upper ejaculatory duct (UED) into whose anterior end the accessory glands and vasa deferentia empty; the funnel characterized by its slit-like lumen; and the lower ejaculatory duct (LED). Anteriorly, the UED has a keyhole-shaped lumen surrounded by a thin intima and highly columnar epithelial cells whose most conspicuous feature is massive aggregations of microtubules. More posteriorly, the UED lumen differentiates into dorsal and ventral chambers, the former having a thick cuticular lining armed with spines. In the hindmost part of the UED, the ventral chamber expands to obliterate the dorsal chamber; its cuticular lining thickens, and conspicuous lateral evaginations develop. The thick cuticle includes 3 distinct layers and on its surface carries numerous spatulate processes. In this region, the epithelial cells develop numerous short microvilli beneath which are many mitochondria. As the funnel is reached, the intima becomes extremely thick, and the epithelial cells lack microvilli and most microtubules. Within the funnel, a new, very distinct form of cuticle appears, which is in “units”, each associated with an epithelial cell and having a rounded epicuticular cap. The new cuticle arises ventrally but rapidly spreads to encircle the entire lumen, at which point the LED is considered to begin. Beneath this new cuticle, the epithelial cells are columnar, have long microvilli, numerous mitochondria in the apical cytoplasm, and rough endoplasmic reticulum basally. Apically, adjacent cells are tightly apposed; however, prominent intercellular channels develop more basally. The ejaculatory duct's features are briefly discussed in terms of its role in spermatophore formation.     

Mesoderm formation in a springtail,Tomocerus ishibashii Yosii (Collembola : Tomoceridae)

The mesoderm formation of Tomocerus ishibashii (Collembola : Tomoceridae) is described. Mesodermal cells are formed after the beginning of the formation of the primary dorsal organ, and originate from the entire region of the embryonic area. After completion of the blastodermic cuticles, cells of mesoderm and ectoderm concentrate towards a ventral midline and form a well-defined 2-layered germ band. The manner of mesoderm formation in the Collembola is similar to that in Diplura and Myriapods, except for the Chilopoda; the mesoderm of the Thysanura s. lat. and Pterygota originates from a localized zone of the embryo. Within the Hexapoda, mesoderm formation is categorized into 2 types: Type 1—unlocalized origin, in the Collembola and Diplura, and Type 2—localized origin, in the Thysanura s. lat. and Pterygota. Types 1 and 2 are thought to be plesiomorphic and apomorphic, respectively.     

Ultrastructure of embryonic envelopes and integument ofOncopeltus fasciatus dallas (Insecta,Heteroptera)

Summary The embryo ofOncopeltus fasciatus forms a typical secondary dorsal organ (SDO). It develops after katatrepsis from the contracting serosa, the cells of which decrease in diameter but increase considerably in height. After 66 h, the SDO represents a protrusion of the serosal epithelium above the head and is then reduced to a disc-shaped formation, which sinks into the yolk, where it disintegrates after 80 h.During its typical expression, between 66 and 78 h, the SDO shows a zonal arrangement of its cell organelles. The nucleus, which is located in the basal cell region, has a very irregular outline and includes several nucleoli and globular inclusion bodies. Rough and smooth ER are well developed around the nucleus and suggest the involvement of the organ in protein secretion as well as in lipid metabolism. Electron-lucent vacuoles and electron-dense granules, sometimes enclosed in the vacuoles, accumulate in the apical cell region, and are obviously extruded into the peripheral (extraembryonic) space. The formation of intercellular clefts and delicate cytoplasmic extensions facing the yolk and microvilli facing the periphery evidence a transporting function of the epithelium. Blisters intercalated in extended junctional complexes between apical cell regions point to the transport of solutes.Because of the similarities of the processes observed in the SDO and in Malpighian tubules of larvae, an excretory function of the SDO is suggested. Final products of yolk and embryo are apparently transported to the extraembryonic space, where they accumulate during embryogenesis.Phylogeny, relationship, and function of the different embryonic glands in Arthropoda (primary and secondary DO and pleuropodia) are discussed.Dedicated to Prof. Dr. B. Scharrer on the occasion of her birthdaySupported by the Deutsche ForschungsgemeinschaftI am grateful to Miss K. Schmidtke and Mrs. M. Ullmann for technical assistance     

Coordinated development in the fly foot: Sequential cuticle secretion

Development of the adult fly foot falls into clearly defined phases of cell division, growth, cuticle secretion and cell death. The pulvillus is composed dorsally of two giant cells and ventrally of thousands of minute tenent cells; the former produce the dorsal footpad cuticle and the latter the thousands of tenent hairs. Cell divisions are still occurring in future tenent cells when increase in size of the cells and in polyteny of the chromosomes is already occurring in the two dorsal cells. Also cell death occurs considerably earlier in the tenent cells, yet the sequential secretion of some six cuticular layers takes place at comparable times in dorsal and ventral cuticles. The cuticular layers formed are, in their order of secretion: ecdysial membrane, cuticulin of the epicuticle, dense exocuticle, homogeneous exocuticle, an intermediate layer, wax of the epicuticle, and an extensive mass of endocuticle. The ecdysial membrane seems to perform an important mechanical role in maintaining the shape of the delicate cytoplasmic projections of the tenent cells, before and during cuticle secretion, and in establishing the cuticular pattern of ridges in the dorsal cuticle. Comparisons are made with trichogen cell cuticle development and with tracheal cuticle. Tracheal, trichogen and dorsal footpad cuticle patterns are compared. Details of giant cell activity provide a working basis for studies of nuclear-cytoplasmic interactions, and the whole system raises many unsolved problems in the general field of cell differentiation and pattern formation.     

Formation of the hindgut cuticular lining during embryonic development of Porcellio scaber (Crustacea,Isopoda)

The hindgut and foregut in terrestrial isopod crustaceans are ectodermal parts of the digestive system and are lined by cuticle, an apical extracellular matrix secreted by epithelial cells. Morphogenesis of the digestive system was reported in previous studies, but differentiation of the gut cuticle was not followed in detail. This study is focused on ultrastructural analyses of hindgut apical matrices and cuticle in selected intramarsupial developmental stages of the terrestrial isopod Porcellio scaber in comparison to adult animals to obtain data on the hindgut cuticular lining differentiation. Our results show that in late embryos of stages 16 and 18 the apical matrix in the hindgut consists of loose material overlaid by a thin intensely ruffled electron dense lamina facing the lumen. The ultrastructural resemblance to the embryonic epidermal matrices described in several arthropods suggests a common principle in chitinous matrix differentiation. The hindgut matrix in the prehatching embryo of stage 19 shows characteristics of the hindgut cuticle, specifically alignment to the apical epithelial surface and a prominent electron dense layer of epicuticle. In the preceding embryonic stage – stage 18 – an electron dense lamina, closely apposed to the apical cell membrane, is evident and is considered as the first epicuticle formation. In marsupial mancae the advanced features of the hindgut cuticle and epithelium are evident: a more prominent epicuticular layer, formation of cuticular spines and an extensive apical labyrinth. In comparison to the hindgut cuticle of adults, the hindgut cuticle of marsupial manca and in particular the electron dense epicuticular layer are much thinner and the difference between cuticle architecture in the anterior chamber and in the papillate region is not yet distinguishable. Differences from the hindgut cuticle in adults imply not fully developed structure and function of the hindgut cuticle in marsupial manca, possibly related also to different environments, as mancae develop in marsupial fluid. Bacteria, evenly distributed within the homogenous electron dense material in the hindgut lumen, were observed only in one specimen of early marsupial manca. The morphological features of gut cuticle renewal are evident in the late marsupial mancae, and are similar to those observed in the exoskeleton.     

The embryonic development of the primitive moth,Neomicropteryx nipponensis Issiki (lepidoptera,micropterygidae): Morphogenesis of the embryo by external observation

The micropterygid moth Neomicropteryx nipponensis belongs to the most primitive suborder Zeugloptera of the Lepidoptera. During embryogenesis the small circular germ disk formed on the ventral egg surface invaginates deeply into the yolk. It finally separates from the egg periphery or rudimentary serosa, and becomes a sac-shaped germ rudiment. Its anterior part later develops into the germ band, while its posterior part is the future amnion. Just before revolution of the embryo, the embryo assumes a completely superficial position beneath the yolk. Neither amnion nor serosa rupture during revolution; by completion of dorsal closure they have been incorporated into the yolk to form the secondary dorsal organ. The formation of the germ rudiment and embryonic membranes in N. nipponensis resembles those of swift moths, Endoclyta (suborder Monotrysia) and of the caddisflies, Stenopsyche (Trichoptera), but differs from those of ditrysian Lepidoptera. The secondary dorsal organ has never been found in any other lepidopteran embryos; however, it is formed in N. nipponensis and in the Trichoptera. The results of the present study strongly support the general phylogenetic views that the Zeugloptera have a close affinity to the Trichoptera.     

Chemical and immunological characterization of proteoglycans of embryonic chick calvaria

We have previously demonstrated in quail embryos grafted on chick yolk sacs the existence of intraembryonic stem cells responsible for definitive hemopoiesis. In order to determine the origin of these cells, we now examine the diffuse hemopoietic processes within the avian embryo's mesoderm. At 4–5 days of incubation in the two species, basophilic cells were found throughout the dorsal mesentery. At 6–8 days these cells became very numerous and built up dense foci at the level of branching of the anterior and posterior cardinal veins. These cells often infiltrated the wall of lymph spaces and channels and were also present in the lumen of blood vessels. Such locations support the interpretation that these basophilic cells represent early stages of hemopoietic differentiation. At 8–10 days, erythropoiesis or granulopoiesis was seen in the foci, which then regressed rapidly. The foci maximal development coincided with the period of colonization of the intraembryonic organ rudiments. In “yolk sac chimeras,” the foci were always constituted by quail cells, indicating their intraembryonic origin. The primordial origin of the intramesodermal cells remains to be determined. A likely source might be the ventral wall of the aorta which appeared to shed cells into the lumen and into the mesentery in the 3-day embryo.     

负子蝽的胚胎发育

负于蝽Sphaerodema rusticus Fabr.的胚胎发育,早期胚带面积大,呈多叶状,明显区分头,颚、胸、腹叶。发育过程需经历胚帝陷入、胚带隆起和胚胎反转三个明显的运动过程。中肠后原基先于前原基形成,并且前、后原基伸展方式不同,使中肠形成不同形态的前、后两部分,两部分的细胞分化亦有差异。神经系统在发育过程中神经节趋于较大程度愈合,腹部的神经节最终愈合为1个复合腹神经节,胸部的神经节也愈合为1个复合胸神经节。本文还总结了胚胎发育时期与卵粒大小的关系,讨论了负子蝽背器官解体的作用。